The genus Micromeria (Lamiaceae), a synoptical update

Willdenowia 38 2008 363 CHRISTIAN BRÄUCHLER, OLOF RYDING & GÜNTHER HEUBL The genus Micromeria (Lamiaceae), a synoptical update Abstract Bräuchler, C., Ryding, O. & Heubl, G.: The genus Micromeria (Lamiaceae), a synoptical update. Willdenowia 38: 363-410. ISSN 0511-9618; 2008 BGBM Berlin-Dahlem. doi:10.3372/wi.38.38202 (available via http://dx.doi.org/) No comprehensive revision of Micromeria is available and uncertainties about the taxonomy of the genus have lasted for a long time. Since the last synopsis many new data from both morphological and molecular genetic studies have been accumulated and, consequently, the number of accepted taxa and the delimitation of the genus have changed considerably. The authors provide a review of recent and unpublished research on the genus, a new circumscription and description of the genus and an updated distribution map. All published Micromeria names are listed with a reference to their current taxonomic position. Names of taxa currently placed in Micromeria are provided with type citations. A new combination, M. cristata subsp. kosaninii, is validated, along with the new name M. longipedunculata for the illegitimate M. parviflora of Reichenbach. The author standard abbreviation E. F. Chapm. is proposed for one of the authors of M. graeca subsp. cypria and 24 names are typified. Taxonomic problems needing special attention in future research are identified. Additional key words: generic circumscription, taxonomy, nomenclature, typification, checklist Introduction The genus Micromeria, distributed from the Macaronesian-Mediterranean region to southern Africa, India and China, was described by Bentham (1829). It belongs to Lamiaceae subfam. Nepetoideae tribe Mentheae subtribe Menthinae and is considered as part of the vaguely defined Satureja complex. To accommodate the apparent morphological diversity many taxonomists have split this complex into several genera, mainly Satureja L., Clinopodium L., Calamintha Mill., Acinos Mill. and Micromeria Benth. (Bentham 1848; Boissier 1879; Ball & Getliffe 1972; Davis 1982; Doroszenko, unpubl. Ph.D. thesis Edinburgh Univ. 1986). Others lumped the group to a single genus Satureja s.l. (Briquet 1896; Brenan 1954; Greuter & al. 1986; Seybold 1988) or Clinopodium (Kuntze 1891: 513-516). These different views on the generic limits of the Satureja complex resulted in considerable taxonomic confusion. Recent studies of the phylogeny and generic status of Satureja s.l. using cladistic analyses of molecular data have contributed a lot to a better understanding of the group. Of particular relevance are papers by Wagstaff & al. (1995), Prather & al. (2002), Trusty & al. (2004) and Bräuchler

364 Bräuchler & al.: Micromeria, a synoptical update & al. (2005). The new data clearly show that Satureja s.l. is not monophyletic. Division of the complex into smaller genera (such as Micromeria) is obviously a more feasible solution than the alternative approach, to achieve monophyly by even further expanding the group to include a number of distinct genera such as Thymus L., Origanum L., Mentha L. and Monarda L. Since the last comprehensive synopsis of Micromeria by Morales (1993), the genus has undergone considerable changes in circumscription and number of accepted taxa. Cantino & Wagstaff (1998), Govaerts (1999) and Harley & Granda (2000) transferred the American species of Micromeria (sect. Xenopoma and sect. Hesperothymus p.p.) to Clinopodium, and their treatments resulted in an updated generic circumscription in Harley & al. (2004). However, recent phylogenetic analyses based on both molecular and morphological/anatomical data (Bräuchler & al. 2005; Bräuchler, unpubl. data) give evidence that Micromeria remains polyphyletic also after the exclusion of the American taxa. Consequently the species of M. sect. Pseudomelissa hadtobetransferred to Clinopodium (Bräuchler & al. 2006; Ryding 2006). The results of the analyses also favour the exclusion of the Old World species of the M. sect. Hesperothymus (from South Africa) and the Malagasy Micromeria species that were omitted by Morales (1993) and placed provisionally in a section Madagascarenses by Doroszenko (unpubl. Ph.D. thesis, Univ. Edinburgh 1986). The latter two groups represent two new genera, one recently described (Bräuchler & al. 2008), the other in preparation (Bräuchler & al. unpubl.). Another still unpublished study by Bräuchler & al., which considers all genera of Menthinae, also shows that M. sect. Cymularia is extraneous in Micromeria (see species enumeration for further details). Exclusion of the above-mentioned groups not only renders Micromeria monophyletic, it also makes the genus more homogeneous, better defined and distinct. Characteristic are leaves with a thickened, almost entire margin. The thickening is caused by a continuous sclerenchymatous ring -vein, and not by a revolute leaf margin as in the Madagascan species hitherto included in Micromeria or in some Clinopodium taxa. This feature is otherwise only shared by very few species of Clinopodium s.l. (e.g., C. nanum, C. corsicum), which, however, differ in calyx characteristics to such an extent that confusion is not likely. The leaves are never conduplicate as in Satureja s.str. or Thymbra. Perfoliate inflorescences, found in, e.g., Mentha, Clinopodium and Micromeria cymuligera, are not found in Micromeria as circumscribed here. Bracteoles are always present. The calyx tube is neither sigmoid nor gibbous at the base (as common in Clinopodium s.l.) and not strongly widening at throat (as in Satureja s.str.). The calyx lobes are usually straight or spreading and not curved. The posterior lip of the corolla is emarginate, curved upwards, not straight; the stamens are not exceeding the corolla lips. As outlined above, four of the six sections recognized by Morales (1993) should be excluded from Micromeria. The remaining two, M. sect. Micromeria and sect. Pineolentia, should be fused. Phylogenetic reconstructions based on molecular data have shown that M. sect. Pineolentia is embedded in a clade of Canarian species of M. sect. Micromeria (Bräuchler & al. 2005; Meimberg & al. 2006). The abandonment of M. sect. Pineolentia is also supported by the fact that the most important diagnostic features (large leaves and large flowers) are shared by some species of M. sect. Micromeria, particularly by the Ethiopian M. unguentaria. These peculiarities could readily be explained by adaptation to the special growth conditions on Gran Canaria. Regarding all available information summarized in the synopsis presented here, it seems not possible to delimit subgroups within Micromeria that are sufficiently divergent to merit recognition at infrageneric level. Due to the great changes in the circumscription of Micromeria, there is a need for a survey on the progress that has been made towards clarifying the species-level taxonomy. A further aim of the synopsis is to provide an updated description of the genus, to give information on typification, accepted names, taxonomically difficult species groups and distribution of the genus. Problems that need to be clarified by more thorough revisional work are additionally emphasized. For taxa that are not part of Micromeria as circumscribed here, but cannot be appropriately accommodated elsewhere, provisional names are indicated. One new name is established, one new combination is made, 24 names are typified.

Willdenowia 38 2008 365 Material and methods The studies conducted by the authors were based on extensive herbarium work, including material of the herbaria B, BM, BOL, BOLO, BP, BR, C, DBN, E, EA, FR, FT, G, GOET, HAL, HBG, HEID, HOH, JE, K, LISC, LZ, M, MA, MO, MPU, MSB, NBG, NU, P, S, SAM, SRGH, STU, TCD, TO, TUB, UPS, W, WAG, WU, Z and ZT (herbarium abbreviations according to Holmgren & Holmgren 1998+) and literature research accompanied by observations of plants both in the wild and in cultivation. Digital images of type specimens cited are found in ALUKA (2008), the Zürcher Herbarien Database (2008), the Virtual Herbaria (2008) and the Virtual Herbarium Berlin (Röpert 2008). Standard abbreviations of authors, periodicals and exsiccata conform to IPNI (2008), BPH online (2008) and IndExs (Triebel & Scholz 2008), respectively. Of great help were also the following digital resources: for literature Bibliotheca Digital CSIC (2008), Botanicus Digital Library (2008), BHL (2008) and Gallica (2008); for collector and author details TL-2 Online (2008); for completion of the names list IPNI (2008) and the World Checklist of Lamiaceae (Govaerts & al. 2008). The work presented here will in turn serve as a revision of Micromeria names in the latter two databases. Micromeria Benth. in Bot. Reg. 15: sub t. 1282. 1829, nom. cons. Type: Micromeria juliana (L.) Benth. ex Rchb., Fl. Germ. Excurs.: 311. 1831 Satureja juliana L., Sp. Pl. 2: 567. 1753. Perennial herbs, subshrubs or shrubs, rarely annual herbs (only a few specimens of M. imbricata from NE tropical Africa), ± aromatic, 2-130 cm tall, with simple hairs and glands. Leaves opposite, petioles usually distinct but short or minute; blade ± leathery, up to 2-30 1-11 mm, often revolute, margin thickened, entire or with a few obscure teeth. Inflorescence thyrsoid or sometimes raceme-like, lax or dense and spike-like; bracts usually similar to the ordinary leaves (sometimes much narrower) but mostly gradually smaller upwards in the inflorescence; cymes 1-many-flowered, often 2 or rarely 3 in the axils of the same node; peduncles minute to 12 mm long, sometimes flat; bracteoles present (also in 1-flowered cymes), small, rather narrow. Calyx 5-lobed, scarcely accrescent, almost actinomorphic to distinctly 2-lipped, 2-11 mm long; tube (12-)13-15-veined with prominent veins, hairy or sometimes glabrous inside near the throat; posterior lip 3-lobed; anterior lip 2-lobed, divided to the base, with the lobes mostly longer and narrower than those of the posterior lip. Corolla strongly 2-lipped, purple, pink violet or white, 3-20 mm long, sometimes shorter and cleistogamous, rarely female; posterior lip emarginate; anterior lip almost flat, 3-lobed with the mid-lobe broader. Stamens 4, ascending under the posterior corolla lip, thecae divaricate, apparently separated on a short connective. Style lobes equal, subequal or sometimes with lower branch slightly longer. Disc rather small, hardly lobed. Nutlets brown or brownish, matt, glabrous or rarely with a single eglandular hair at apex (in M. imbricata), 0.6-1.5 0.3-0.8 mm, 1.15-4 times longer than broad, apex acuminate, acute, subacute or rounded to obtuse, with a ± distinct areole at the scar, producing mucilage when wet. 2n = 20(?), 26, 30, 48, 60. No sections accepted within the genus. As circumscribed here, Micromeria includes about 54 accepted species with 32 subspecies and 13 varieties. The number will probably change in the near future since on the one hand some new species are to be described while on the other hand at least 15 of the species are doubtful concerning their distinctness from closely related taxa and might prove conspecific. The infraspecific classification in many parts is in need of a thorough revision. The name Micromeria has been conserved against Xenopoma Willd. in Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin 5: 399. 1811 and Zygis Desv.inW.Hamilton, Prodr. Pl. Ind. Occid.: 46. 1825 (Babu 1969), both then monotypic New World genera. The corresponding types represent one and the same species, today included in Clinopodium (C.

366 Bräuchler & al.: Micromeria, a synoptical update Fig. 1. Distribution map of Micromeria s.str. vimineum (L.) Kuntze) and thus are no longer regarded as congeneric with the species here included in Micromeria. Distribution Mediterranean area, Portugal, Madeira, Canary Islands, Cape Verde Islands, E Nigeria, Equatorial Guinea (Bioco), Cameroon, Tschad (Tibesti), Sudan, Ethiopia, Eritrea, Djibouti, Somalia, D. R. Congo, Rwanda, Burundi, Uganda, Kenya, Tanzania, Angola (Huila Plateau), E Zambia, Malawi, Mozambique (Manica Prov.), South Africa, Lesotho, SW Saudi Arabia, Yemen, Oman, Iraq (Kurdistan), Iran, Afghanistan, N Pakistan, Nepal, Bhutan, India, Myanmar, China (Yunnan and Guizhou) (Fig.1). Reports from S Algeria and N Niger are doubtful and need confirmation by recollecting. There are uncertainties about the distribution area in Myanmar and China. Rearching from sea level to more than 2000 m in the Mediterranean and Macaronesia; 400-4500(?-4800 m) in tropical to subtropical Africa and Asia; mostly occurring in moderately dry areas with a rather long dry season. List of published names All names published in Micromeria at the rank of species or below are recorded in alphabetical order. Currently accepted names are marked in bold, invalidly published names are put in square brackets. The records of synonyms contain a cross reference to the currently accepted name. The basionyms are given. Infraspecific taxa are listed alphabetically under the species entry irrespective of rank; combinations of the same basionym at different ranks are united under one entry. Autonyms are only included where infraspecific taxa are accepted. Types belonging to Micromeria names are listed, but types of names that are referred to other genera have been omitted.

Willdenowia 38 2008 367 The type citation is given under the basionyms, or if the basionym was not described in Micromeria, under the currently accepted Micromeria name. Type citations are made to the best of our knowledge, and where information was lacking this is explicitly stated. Herbarium barcode numbers (where present) are cited for type specimens that we have only seen as digital images, those from ALUKA (2008) are additionally marked with an asterisk (*). In some cases digital images have been provided directly by the curators of the corresponding herbaria (BOLO, G, STU, TO). A regularly updated version of this list will be made available at the homepage of the first author s institution. Micromeria abyssinica Hochst. [in Flora 24, Intell. 2: 23. 1841, nom. nud.] ex Benth. in Candolle, Prodr. 12: 224. 1848 Clinopodium abyssinicum (Hochst. ex Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. acropolitana Halácsy ex Maire & Petitmengin in Bull. Soc. Sci. Nancy, ser. 3, 9: 421. May 1908 & Halácsy, Consp. Fl. Graec., Suppl. 1: 87. Jun 1908. Lectotype (designated here by Bräuchler): [Greece] In Acropoli Athenarum, 30.8.1906, Maire & Petitmengin, Mission Botanique en Orient no. 1073 (WU [herb. Halácsy]!). =? M. microphylla (d Urv.) Benth. Note. The NCY syntype could not be traced so far (C. Drechsler, pers. comm.), the WU syntype after several fruitless efforts, however, was found by H. Rainer and W. Till stored under the unpublished name Micromeria athenae. This specimen contains a handwritten diagnosis by Halácsy and is here designated as lectotype. Nowhere in the treatment containing the protologue or one of its prior parts published in earlier volumes of the same journal it is stated that Halácsy contributed to the descriptions in any way. The authorship therefore has to be attributed to Maire & Petitmengin, and this publications predates Halácsy s own publication of the species by one month. The status of this taxon is not settled, it seems very similar to the type of M. sphaciotica Boiss. & Heldr. ex Benth. and thus might not be specifically distinct from M. microphylla (d Urv.) Benth. (see under the respective name entries below). [M. aetnensis Lacaita in Lojacono-Pojero, Fl. Sicul. 2(2): 211. 1907, pro syn.] M. graeca (L.) Benth. ex Rchb. [M. aetnensis var. elata Lacaita in Lojacono-Pojero, Fl. Sicul. 2(2): 211. 1907, pro syn.] M. graeca (L.) Benth. ex Rchb. [M. aetnensis var. humilis Lacaita in Lojacono-Pojero, Fl. Sicul. 2(2): 211. 1907, pro syn.] M. graeca (L.) Benth. ex Rchb. M. affinis Hook. f. in J. Bot. 6: 274. 1847 = Mentha diemenica Spreng., Syst. Veg. 2: 724. 1825. M. afghanica Freitag in Notes Roy. Bot. Gard. Edinburgh 31: 353. 1972 = Gontscharovia popovii (B. Fedtsch. & Gontsch.) Boriss. in Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR 15: 321. 1953. M. albanica (Griseb. ex K. Mal1) Sili6, Monogr. Satureja Fl. Jugosl.: 202. 1979 Satureja albanica Griseb.exK.Mal1 in Bull. Inst. Jard. Bot. Univ. Belgrade 1-3: 229-230. 1930 = Clinopodium serpyllifolium (M. Bieb.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. alpestris Urb. in Repert. Spec. Nov. Regni Veg. 16: 143. 1919 Clinopodium alpestre (Urb.) Harley in Kew Bull. 55: 919. 2000. M. alternipilosa K. Koch in Linnaea 19: 25. 1846 = Satureja spicigera (K. Koch) Boiss., Fl. Orient. 4: 566. 1879. M. amana Rech. f. in Svensk. Bot. Tidsskr. 43: 42. Mar 1949. Holotype: [Turkey] Syria borealis: Mons Amanus, mont. de Düldül, 1500-2000 m, 8.1911, Haradjan 3887 (G; isotypes: E!, W!). M. cremnophila subsp. amana (Rech. f.) P. H. Davis.

368 Bräuchler & al.: Micromeria, a synoptical update Note. The W isotype has no original label and the collection date apparently was retyped erroneously as avril instead of août as is mentioned in the protologue and written on the label of the E isotype. M. approximata (Biv.) Rchb., Fl. Germ. Excurs.: 859. 1832, nom. illeg. Satureja approximata Biv., Stirp. Rar. Sicilia 4: 13. 1818, nom. illeg. Satureja fasciculata Raf., Précis Découv. Somiol.: 39. 1814 = M. graeca subsp. fruticulosa (Bertol.) Guinea Note. Both Bivona-Bernardi and Reichenbach included the older valid name Satureja fasciculata Raf. in the synonymy rendering their names illegitimate. M. approximata subsp. barceloi (Willk.) Nyman, Consp. Fl. Eur.: 590. 1881, nom. illeg. M. approximata [unranked] barceloi (Willk.) Gand., Nov. Consp. Fl. Eur.: 397. 1910, nom. illeg. M. barceloi Willk. = M. inodora (Desf.) Benth. Note. Gandoger (1910) listed several names as infraspecific taxa without indication of rank ( sous-espèces, races ou formes ). His names therefore are validly published but inoperative in questions of priority except for homonymy (see McNeill & al. 2006: Art. 35.3 & 53.4). This combination, however, is illegitimate, because it includes the legitimate name barceloi under the illegitimate approximata. [M. approximata var. cymulosa Lojac., Fl. Sicul. 2(2): t. XVIII. 1907, nom. inval.] M. graeca (L.) Benth. ex Rchb. Note. This name applies to an illustration not accompanied by an analysis (McNeill & al. 2006: Art. 42.4) and therefore is not validly published. M. arganietorum (Emb.) R. Morales in Anales Jard. Bot. Madrid 48: 153. 1991 Satureja arganietorum Emb. in Bull. Soc. Sci. Nat. Maroc 15: 182. 1936. Holotype: [Morocco] Anti Atlas occidental: Falaises siliceuses dominant le gué de Taderrast sur l oued Massa, 200 m, 30.4. 1934, Emberger s.n. (MPU-006094*!). M. arkansana (Nutt.) Benth., Labiat. Gen. Spec.: 730. 1835 Hedeoma arkansana Nutt. in Trans. Amer. Philos. Soc., ser. 2, 5: 186. 1834 = Clinopodium glabrum (Nutt.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. australis (R. Br.) Benth., Labiat. Gen. Spec.: 380. 1834 Mentha australis R. Br., Prodr. Fl. Nov. Holland.: 505. 1810. M. bahamensis Shinners in Sida 1: 96. 1962 = Clinopodium brownei (Sw.) Kuntze, Revis. Gen. Pl. 2: 514. 1891. M. balcanica Velen., Fl. Bulg. Suppl. I: 235. 1898 = Clinopodium frivaldszkyanum (Degen) Bräuchler & Heubl in Taxon 55: 979. 2006. Note. Doroszenko (unpubl. Ph.D. thesis, Univ. Edinburgh 1986) includes this name in the synonymy of M. frivaldszkyana, which is followed here. A final decision on the synonymy of Clinopodium dalmaticum (Benth.) Bräuchler & Heubl, C. frivaldszkyanum (Degen) Bräuchler & Heubl and C. pulegium (Rochel) Bräuchler cannot be made without a thorough revision of the group, which is in progress but beyond the scope of this manuscript. M. balcanica var. moesica Urumoff in Oesterr. Bot. Z. 49: 203. Jun 1899 = Clinopodium frivaldszkyanum (Deg.) Bräuchler & Heubl in Taxon 55: 979. 2006. Note. See note under M. balcanica. M. barbata C. A. Mey. in Fischer & Meyer, Index Seminum (St. Petersburg) 8: 67. 1842 = Clinopodium douglasii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. barbata Boiss. & Kotschy in Boissier, Diagn. Pl. Orient., ser. 2, 4: 14. 1859, non C. A. Mey. 1842 Clinopodium serpyllifolium subsp. barbatum (P. H. Davis) Bräuchler in Taxon 55: 980. 2006.

Willdenowia 38 2008 369 M. barceloi Willk. in Oesterr. Bot. Z. 25: 111. 1875. Lectotype (designated here by Bräuchler): [Spain] Baleares, Mallorca colina calcarea al O. de Palma, 14.11.1873, Willkomm s.n. (MA!). = M. inodora (Desf.) Benth. [M. barceloi var. africana Batt. in Bull. Soc. Bot. France 44: 323. 1897, nom. inval.] M. inodora (Desf.) Benth. Note. This variety was described from Algeria as being simply more robust than the typical variety. Given this apparent lack of a diagnosis, we consider this name not validly published. M. barosma (W. W. Sm.) Hand.-Mazz., Symb. Sin. 7: 932. 1936 Calamintha barosma W. W. Sm. in Notes Roy. Bot. Gard. Edinburgh 9: 88. 1916 Clinopodium barosmum (W.W. Sm.) Bräuchler & Heubl in Taxon 55: 978. 2006. M. benthamii Webb & Berthel., Hist. Nat. Iles Canaries 3: 77. 1845 [as benthami ]. Lectotype (designated by Pérez 1978: 122): [Spain, Canary Islands] In rupestribus altioribus Magna Canariae prosertum in jugo Montium Saucillo, Barker-Webb (FI [lower individual]). M. biflora (Buch.-Ham. ex D. Don) Benth., Labiat. Gen. Spec.: 378. 1834 Thymus biflorus Buch.-Ham. ex D. Don, Prodr. Fl. Nepal.: 112. 1825. Type: [Nepal] Ad Suembu Nepaliae superioris, Hamilton s.n. (not traced, probably lost). Note. Perhaps better included in M. imbricata or considered as a variety of this species. The type could not be traced at BM, E or K. [M. biflora subsp. arabica K. H. Walther in Collenette, Illustr. Guide Wildfl. Saudi Arab.: 273. 1985 & in Collenette, Wildfl. Saudi Arab.: 452. 1999 & in Al-Rehaily in Pakistan J. Biol. Sci. 9: 2726-2728. 2006, nom. nud.] M. imbricata (Forssk.) C. Chr. var. imbricata M. biflora var. cinereotomentosa (A. Rich.) Chiov. in Nuov. Giorn. Bot. Ital., ser. 2, 26: 163. 1919 M. ovata var. cinereotomentosa A. Rich. = M. imbricata (Forssk.) C. Chr. var. imbricata M. biflora f. filiformis (Aiton) Knoche, Fl. Baléar. 2: 348. 1922, nom. illeg. M. filiformis (Aiton) Benth. Note. Knoche included the indirect reference M. filiformis Benth. to the older name Thymus filiformis Aiton, when establishing filiformis as infraspecific taxon under M. biflora (Buch.-Ham. ex D. Don) Benth., making his new combination illegitimate. M. biflora var. hirsuta Fiori in Nuov. Giorn. Bot. Ital., ser. 2, 20: 371. 1913. Holotype: [Eritrea] Acchelè-Guzai, tra Mai-Harazat ed Halai, Fiori 1570 (not traced). = M. imbricata (Forssk.) C. Chr. var. imbricata M. biflora var. indica Elly Walther & K. H. Walther in Mitt. Thüring. Bot. Ges. 1(4): 6. 1957. Holotype: [India, Prov. Medes] Cornon, Coonoor (HBG!). = M. biflora (Buch.-Ham. ex D. Don) Benth. M. biflora var. punctata (Benth.) Fiori in Nuov. Giorn. Bot. Ital., ser. 2, 20: 371. 1913 M. punctata Benth. = M. imbricata (Forssk.) C. Chr. var. imbricata M. biflora var. rhodesiaca Elly Walther & K. H. Walther in Mitt. Thüring. Bot. Ges. 1(4): 7. 1957. Holotype: [Malawi] Mt Mlanje, Adamson 368 (K!). M. imbricata var. rhodesiaca (Elly Walther & K. H. Walther) Ryding M. biflora var. rodriguezii (Freyn & Janka) Knoche, Fl. Baléar. 2: 346. 1922 M. rodriguezii Freyn & Janka M. filiformis subsp. rodriguezii (Freyn & Janka) Bonafè M. biflora var. typica Elly Walther & K. H. Walther in Mitt. Thüring. Bot. Ges. 1(4): 6. 1957 [nom. inval.?]. Holotype: [India] Kamoun, Wallich (K!). = M. biflora (Buch.-Ham. ex D. Don) Benth.

370 Bräuchler & al.: Micromeria, a synoptical update Note. Despite not explicitly including the type of Micromeria biflora, this variety name could be regarded as not validly published (McNeill & al. 2006: Art. 24.3). The protologue, however, includes a type citation (not the type of the species), a diagnosis and a status designation as var. nov.. M. biflora var. villosa Elly Walther & K. H. Walther in Mitt. Thüring. Bot. Ges. 1(4): 7. 1957. Holotype: [Kenya] Klinangop, Dale 2965 (K!). M. imbricata var. villosa (Elly Walther & K. H. Walther) Ryding M. boliviana Benth., Labiat. Gen. Spec.: 731. 1835 Clinopodium bolivianum (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. boliviana var. angustifolia Wedd., Chlor. Andina 2: 150. 1860 = Clinopodium bolivianum (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. bonariensis (Ten.) C. A. Mey. in Fischer & Meyer, Index Seminum (St. Petersburg) 10: 56. 1845 Thymus bonariensis Ten. in Index Seminum (Napoli) 1839: 12. 1839 = Hedeoma multiflora Benth., Labiat. Gen. Spec.: 367. 1834. M. bourgaeana Webb ex Bolle in Bonplandia (Hannover) 8: 283. 1860. Holotype: [Spain, Canary Islands] Hab. in Canaria Magna in rupestribus loci dicti Arecife, rara, Bourgeau [Bourgeau, Pl. Canar. 1845-46, 508] (B ; isotypes: K!, P!, TCD!, W!, WU!, Z!). = M. tenuis subsp. linkii (Webb & Berthel.) P. Pérez M. bracteolata (Nutt.) Benth., Labiat. Gen. Spec.: 371. 1834 Hedeoma bracteolata Nutt., Gen. N. Amer. Pl., Addend.: 4. 1818 = Conradina sp. Note. Synonymy follows Epling & Stewart (1939: 46). M. brevicalyx (Epling) R. Morales in Bot. Complut. 18: 164. 1993 Satureja brevicalyx Epling in Ann. Missouri Bot. Gard. 14: 82. 1927 Clinopodium brevicalyx (Epling) Harley & A. Granda in Kew Bull. 55: 919. 2000. M. brivesii Batt. in Bull. Soc. Hist. Nat. Afrique N. 13: 69. 1922. Type: [Algeria] Près de l Oued Amismiz, 4.11.1909, Brives s.n. (not traced, MPU?). Note. Some other Brives types are present at MPU, but this one is not included in ALUKA (2008). According to Stafleu & Cowan (1976) part of Battandier s types are present at AL, but more recent information (P. Schäfer, pers. comm.) indicate these collections either as transferred to MPU or lost. M. browiczii Ziel. & Kit Tan in Polish Bot. J. 46: 31. 2001. Holotype: Greece, Ionian Islands. Nomos and eparchia of Zakinthou: Zakinthos, between Agalas and the promontory of Ketinaria, calcareous rocky slope, c. 100 m, 25.5.1988, Boratysoki, Browicz, Tomlik & Zielisoki 765 (KOR; isotypes: ATH, herb. Kit Tan!). M. brownei (Sw.) Benth., Labiat. Gen. Spec.: 372. 1834 Thymus brownei Sw., Prodr. Descr. Veg.: 89. 1788 Clinopodium brownei (Sw.) Kuntze, Revis. Gen. Pl. 2: 514. 1891. Note. Sometimes erroneously listed as M. brownii. M. brownei var. ludens Shinners in Sida 1: 96. 1962 = Clinopodium brownei (Sw.) Kuntze, Revis. Gen. Pl. 2: 514. 1891. M. brownei var. pilosiuscula A. Gray, Syn. Fl. N. Amer. 2(1): 359. 1878 = Clinopodium brownei (Sw.) Kuntze, Revis. Gen. Pl. 2: 514. 1891. M. bucheri P. Wilson in J. New York Bot. Gard. 23: 93. 1922 Clinopodium bucheri (P. Wilson) Harley in Kew Bull. 55: 920. 2000. M. buchii Webb & Berthel., Hist. Nat. Iles Canaries 3: t. 161. Jun 1849 - Mar 1850, nom. illeg. Type: forecited illustration (apparently based on a specimen at FI labelled Micromeria buchii Webb ). M. tenuis subsp. linkii (Webb & Berthel.) P. Pérez

Willdenowia 38 2008 371 Note. Table 161 represents an illustration with an analysis (McNeill & al. 2006: Art. 42.4) and M. buchii would thus have been validly published. The corresponding illustration legend and species description in the text volume of the same work were, however, published at least four years in advance (l.c.: 73. Apr 1845). Since both refer to M. linkii, M. buchii is to be regarded as illegitimate. In addition, Peréz (1978: 138) designated a specimen at FI labelled Micromeria buchii as lectotype for M. linkii. Dates for the Webb names are taken from Stearn (1937). M. bulgarica (Velen.) Vandas in Magyar Bot. Lapok 4 (8-11): 267. 1905 M. origanifolia subsp. bulgarica Velen. = Clinopodium dalmaticum (Benth.) Bräuchler & Heubl in Taxon 55: 979. 2006. Note. See comments under M. balcanica Velen. The citation M. bulgarica (Velen.) Hayek in Repert. Spec. Nov. Regni Veg. Beih. 30(2): 323. 1929, is incorrect, because the latter author refers in a footnote to Vandas name. M. byzantina (K. Koch) Walp., Ann. Bot. Syst. 3: 251. 1852 Calamintha byzantina K. Koch in Linnaea 21: 672. 1849 = Clinopodium nepeta subsp. glandulosum (Req.) Govaerts, World Checklist Seed Pl. 3(1): 18. 1999. M. calaminthoides Lojac., Fl. Sicul. 2(2): 212. 1907. Syntypes: [Italy, Sicily] Rupi di M. Pellegrino (?) 3.1825, Tineo, inherb.pan.&herb.tod.subm. graeca longiflora (not traced). = M. graeca (L.) Benth. ex Rchb. Note. ManyoftheMicromeria names published by Lojacono-Pojero (1907) without doubt are to be placed in synonymy to M. graeca. Judging from the diagnoses and the accompanying illustrations it is, however, often not possible to assign the taxa to one of the currently accepted subspecies without revising the whole group. Types of Lojacono s names have not been traced for this paper. His herbarium according to Stafleu & Cowan (1981) is integrated in PAL. M. calaminthoides var. contracta Lojac., Fl. Sicul. 2(2): 213. 1907. Syntypes: [Italy, Sicily] Palermo alla Scala di Maseddo, Tineo, Monti di Castellammare, Gibilrossa, Busambra (not traced). = M. graeca (L.) Benth. ex Rchb. M. calaminthoides var. elongata Lojac., Fl. Sicul. 2(2): 213. 1907. Type: [Italy, Sicily] Pizzuta ( herb. Pan., not traced). = M. graeca (L.) Benth. ex Rchb. M. calostachya Rech. f. in Denkschr. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 105: 122. 1943 Thymbra calostachya (Rech. f.) Rech. f. in Kulturpflanze, Beih. 3: 64. 1962. [M. calvertii Boiss. in Fl. Orient. 4: 571. 1879, pro syn.] M. elliptica K. Koch M. canescens (Guss.) Benth., Labiat. Gen. Spec.: 376. 1834 Satureja canescens Guss., Pl. Rar.: 228, t. 42. 1826. Syntypes: [Italy] In collibus maritimis Japygiae: Taranto, Punta di Cutaro, Gallipoli, Torricella, Otranto, Gussone (NAP?). = M. graeca (L.) Benth. ex Rchb. Note. This name is accepted at species rank by Pignatti (1982) and with doubt placed in synonymy to Satureja graeca subsp. garganica by Briquet (1895: 421). Doroszenko (unpubl. Ph.D. thesis, Univ. Edinburgh 1986) places it in synonymy to M. graeca subsp. longiflora. Regarding Gussone s description and illustration, the latter might be more appropriate. M. canescens (K. Koch) Walp., Ann. Bot. Syst. 3: 251. 1852, non Benth. 1834 Calamintha canescens K. Koch in Linnaea 21: 672. 1849 = Clinopodium nepeta (L.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. capitellata Benth. in Candolle, Prodr. 12: 218. 1848 Clinopodium capitellatum (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. carica P. H. Davis in Kew Bull. 1949: 109. 1949 Clinopodium caricum (P.H.Davis)Bräuchler & Heubl in Taxon 55: 978. 2006.

372 Bräuchler & al.: Micromeria, a synoptical update M. carminea P. H. Davis in Kew Bull. 1949: 400. 1949. Holotype: Turkey, Denizli: d. Acipayam, Boz Da. above Geyran yayla, 1800-1900 m, cervices of limestone rock with Globularia dumulosa, flowers almost pure carmine, 16.7.1947, Davis 13403 (K!; isotypes: E!, W!). M. cristata subsp. carminea (P.H.Davis)P.H.Davis M. carpatha Rech. f. in Phyton 1: 208. 1949. Lectotype (designated here by Bräuchler): [Greece] Karpathos, 5.6.1886, Forsyth Major 165a [protologue: Vrondi, FM. 165a in hb. Barbey ] (G-00087102!). = M. microphylla (d Urv.) Benth. Note. This name is sometimes placed in synonymy to M. sphaciotica Boiss. & Heldr. ex Benth., which in turn mostly has been included in synonymy to M. microphylla (d Urv.) Benth. (e.g., Boissier 1879; Sili6 1979; Doroszenko, unpubl. Ph.D. thesis, Univ. Edinburgh 1986). This opinion is accepted here. The second syntype cited is Karpathos, Rechinger 8249 (W). M. chamissonis (Benth.) Greene, Man. Bot. San Francisco: 289. 1894 Thymus chamissonis Benth. in Linnaea 6: 80. 1831 = Clinopodium douglasii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. [M. chanonis Muschl. in Just s Bot. Jahresber. 36(3): 607. 1912, erroneous for M. chamissonis (Benth.) Greene]. Clinopodium douglasii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. Note. When citing an article of Power & Salway (1908), Muschler mistranslated the original title Chemical examination of Micromeria chamissonis to Chemische Untersuchung von Micromeria chanonis (Verba buena), the name thus merely represents a type error. M. chionistrae Meikle in Ann. Mus. Goulandris 6: 92. 1983. Holotype: Cyprus, Phini, 1000 m, in cracks of bare rock, 6.6.1939, Kennedy 1495 (K!). M. cilicica Hausskn. ex P. H. Davis in Kew Bull. 1949: 109. 1949 Clinopodium cilicicum (Hausskn. ex P. H. Davis) Bräuchler & Heubl in Taxon 55: 978. 2006. [M. cinerea Doroszenko in Ryding in Bot. J. Linn. Soc. 155: 438. 2007, pro syn.] M. imbricata (Forssk.) C. Chr. var. imbricata Note. This name was given to a new species by Doroszenko s (unpubl. Ph.D. thesis, Univ. Edinburgh 1986), but is neither effectively nor validly published. Ryding (2007), however, mentions this name under M. imbricata var. imbricata. It therefore is included here to complete the list of names found in literature available to us. M. compacta (Killick) R. Morales in Bot. Complut. 18: 164. 1993 Satureja compacta Killick in Bothalia 7: 437. 1961 Killickia compacta (Killick) Bräuchler, Doroszenko & Heubl in Bot. J. Linn. Soc. 157: 584. 2008. Note. One of three species endemic to the South Afrikan Drakensberg. All three have previously been included in Satureja or Micromeria sect. Hesperothymus. While New World members of the latter section have been transferred to Clinopodium, these three species remained in Micromeria and recently have been placed in a genus of their own named Killickia (Bräuchler & al. 2008). M. conferta (Coss. & Daveau) Stefani in Boll. Reale Orto Bot. Palermo 11: 148. 1912 M. juliana var. conferta Coss. & Daveau Note. Obviously this was intended to be a new combination for M. juliana var. conferta Coss. & Daveau and according to Art. 33.2 and 33.3 of the Vienna Code (McNeill & al. 2006) needs to be treated as such. The reference to the basionym, however, is very cryptic ( Micromeria conferta Cass. apparently a miss-citation of Coss. & Daveau).The fact that only one specimen is cited and a description (in Italian) is given obscures the new combination even more. This species is included under M. juliana by Siddiqi (1985). M. congesta Boiss. & Hausskn. in Boissier, Fl. Orient. 4: 575. 1879 Clinopodium congestum (Boiss. & Hausskn.) Kuntze, Revis. Gen. Pl. 2: 515. 1891.

Willdenowia 38 2008 373 M. consentina (Ten.) N. Terracc. in Nuov. Giorn. Bot. 5: 227. 1873 [as M. cosentina ] Satureja consentina Ten. M. graeca subsp. consentina (Ten.) Guinea M. consentina var. aeolica Lojac., Fl. Sicul. 2(2): 210. 1907. Syntypes: [Italy] Isole Eolie: colli vulcanici: Panaria, Lojacono-Pojero, Salina a S Marina, Lojacono-Pojero, Alicuari Tineo, Lipari,Lojacono-Pojero (not traced). = M. graeca subsp. consentina (Ten.) Guinea Note. See note under M. calaminthoides. M. consentina var. chamaepitys Lojac., Fl. Sicul. 2(2): 210. 1907. Syntypes: [Italy, Sicily] Ad rupes graniticas v. schistosas in Sicilia sept. Capo d Orlando, Milazzo, Antennamare, Gussone, Prov. Messina, Gussone (not traced). = M. graeca subsp. consentina (Ten.) Guinea M. consentina var. glabrata (Guss.) Strobl in Flora 68: 369. 1885 Satureja consentina var. glabrata Guss., Fl. Sicul. Syn. 2: 88. 1844. Syntypes: Palermo a S. Maria a Gesu, ed alla Pizzuta, Catania, Aci, Giarre, Scaletta, Capo S. Alessio, Taormina, Messina, Capo di Calava, Capo Tindaro (NAP?). = M. graeca subsp. consentina (Ten.) Guinea [M. consentina var. glabrescens Lojac., Fl. Sicul. 2(2): 210. 1907, pro syn.] M. graeca subsp. consentina (Ten.) Guinea M. consentina var. minutissima Lojac., Fl. Sicul. 2(2): 210. 1907. Type: [Italy, Sicily] Ad rupes vulcanicas apricas calidissimas in coespites late pulvinatos pulchre floriferos, Ins. Alicuri, 1.8.1905 (NAP?). = M. graeca subsp. consentina (Ten.) Guinea M. consentina var. pseudotodari Lacaita ex Lojac., Fl. Sicul. 2(2): 210. 1907. Syntypes: [Italy, Sicily] rupi primitive: Castanea, Todaro, Capo d Orlando, Fiumara di Zappulla, Sic. Nord.-Orient, Todaro, Capo Tindaro, Gussone, Capo Calava, Gussone, Vetta dell Antennamare, Gussone (not traced). = M. graeca subsp. consentina (Ten.) Guinea M. consentina var. rossii Lojac., Fl. Sicul. 2(2): 210. 1907 [as rossi ]. Syntypes: [Italy, Sicily] Aetna, Rossi, Nicolosi, Tineo, Torn., Catania Chiuse Torn., Tineo, Luglio (not traced). = M. graeca subsp. consentina (Ten.) Guinea M. cordata (Moris ex Bertol.) Moris, Diag. Stirp. Sard. Nov.: 2. Mar 1857 Satureja cordata Moris ex Bertol., Fl. Ital. 10: 519. Jan-Feb 1857. Lectotype (designated here by Bräuchler): [Italy, Sardinia] Moris (BOLO [photo!]). Note. This species is included as subspecies in M. filiformis (Pignatti 1977) or in M. microphylla (Doroszenko unpubl. Ph.D. thesis, Univ. Edinburgh 1986). Further studies are needed to examine the delimitation between the two taxa, even more so since obvious morphological and ecological affinities with M. marginata exist. For the time being we think it is best to follow Valsecchi (1978) and treat it as a distinct species. The latter author has designated a lectotype for M. cordata Moris from TO ( M. cordata Moris, nelle fissuri delle rupi, Monti di Oliena, Giugno 1852, photo!). Bertoloni s name, however, has priority and there is no evidence that the Moris collection received by him was a duplicate of the lectotype, consequently the typification by Valsecchi is rejected here. The only specimen preserved at BOLO and annotated as Satureja cordata Moris by Bertoloni has no locality on the label but a note indicating it as sent by Moris in 1856, which means prior to publication of the species description. This is the only collection unambiguously representing original material and therefore is chosen as lectotype here. M. cordata [unranked] allionii (Zumagl.) Gand., Nov. Consp. Fl. Eur.: 397. 1910 Thymus allionii Zumagl., Fl. Pedem. 1: 226. 1859 Thymus piperella All., Fl. Pedem. 1: 21, t. 37, fig. 3. 1785, non L. 1753 M. marginata (Sm.) Chater Note. See note under M. approximata subsp. barceloi. M. corsica (Pers.) H. Lév., Dict. Invent. Fl. Franc.: 22. 1916 Thymus corsicus Pers., Syn. Pl. 2: 131. 1806 Acinos corsicus (Pers.) Getliffe in Bot. J. Linn. Soc. 65: 263. 1972.

374 Bräuchler & al.: Micromeria, a synoptical update M. cremnophila Boiss. & Heldr. in Boissier, Fl. Orient. 4: 570. 1879. Holotype: [Greece] Ad rupes verticales regionis abietinae Parnassi ad Gourna 4000'-4500', Heldreich (G-BOIS). M. cremnophila subsp. amana (Rech. f.) P. H. Davis in Notes Roy. Bot. Gard. Edinburgh 38: 40. 1980 M. amana Rech. f. M. cremnophila subsp. anatolica P. H. Davis in Notes Roy. Bot. Gard. Edinburgh 38: 40. 1980. Holotype: Turkey B9 Van: 5 km N of Certak, in cervices of boulders, 25.7.1954, Davis 23258 & Polunin (E!; isotype: K!). M. cremnophila Boiss. & Heldr. subsp. cremnophila M. cremnophila var. glandulosa Micevski in Prilozi Oddel. Biol. Med. Nauki, Makedonska Akad. Nauk. Umet. 23: 17. 2002. Holotype: [Macedonia] In fauce ad flumen Radika inter pagos Trnica et Volkovija, in rupibus calcareis 800 m, 23.8.1986, Micevski (SKO). = M. cremnophila Boiss. & Heldr. subsp. cremnophila M. cremnophila var. thessala Formánek in Verh. Naturf. Vereins Brünn 35: 142 & 183. 1896. Type: [Greece] Thessalia, Pindus: Oxya Despoti, untere Lage 210-400 m, Kastri et in sylvaticis Metochi Vavlavi, pr. Vlachava, 1896, Formánek (not traced). = M. cremnophila Boiss. & Heldr. subsp. cremnophila M. cristata (Hampe) Griseb., Spicil. Fl. Rumel. 2: 122. 1844 Thymus cristatus Hampe in Flora 20: 233. 1837. Lectotype (designated here by Bräuchler): [Bulgaria] Rumelia, [1835] Frivaldszky (BP-334973!). Note. This plant has been distributed by Frivaldszky (1836) as part of his Centuria plantarum exsiccatarum anno 1835 in Turcia-europaea collectarum under the name Thymus inodorus Desf. It was collected in Rumelia (probably in the Rhodopian mountains) by Füle, Hinke or Manolesko (all then collectors sent out by Frivaldszky; Degen 1896). Hampe (1837) subsequently identified the plant as representing a new species, which he described as T. cristatus. Since the whereabouts of his herbarium of phanerogams are uncertain, a specimen of Frivaldsky s Centuria traced at BP is designated as lectotype here. The plants on this sheet represents part of the original material, probably an isotype (though neither annotated by Hampe nor labelled with his name, but Frivaldszky s original determination instead). No duplicates have been traced so far, but might be present in numerous herbaria (e.g., G, K, P, W). M. cristata f. albiflora Sili6 in Glasn. Zemaljsk. Muz. Bosne Hercegovine Sarajevu Prir. Nauke 13: 116. 1975. Holotype: [Serbia] Jelasni6ka klisura prope Nis, in valle fluminis, 22.8.1953, Lindtner (BEO). = M. cristata (Hampe) Griseb. subsp. cristata M. cristata f. canescens Vandas in Rel. Form.: 483. 1909 M. cristata subf. canescens (Vandas) Hayek, Prodr. Fl. Penins. Balcan. 2: 319. 1929 M. cristata var. canescens (Vandas) Sili6 in Glasn. Zemaljsk. Muz. Bosne Hercegovine Sarajevu Prir. Nauke 13: 116. 1975. Syntypes: [Macedonia] Plantae e Flora et Hadzibarica pl., Formánek (not traced). = M. cristata (Hampe) Griseb. subsp. cristata M. cristata subsp. carminea (P. H. Davis) P. H. Davis in Notes Roy. Bot. Gard. Edinburgh 21: 65. 1952 M. carminea P. H. Davis M. cristata (Hampe) Griseb. subsp. cristata M. cristata var. glandulosa Micevski in Prilozi Oddel. Biol. Med. Nauki, Makedonska Akad. Nauk. Umet. 23: 12. 2002. Holotype: [Macedonia] Fauces Demir Kapija, 9.6.1972, Micevski (SKO). = M. cristata (Hampe) Griseb. subsp. cristata Micromeria cristata subsp. kosaninii (Sili6) Bräuchler & Govaerts, comb. nov. Micromeria kosaninii Sili6, Monogr. Satureja Fl. Jugosl.: 234. 1979. Holotype: [Macedonia] Galicica, Poljce, c. 1600 m, solo calcareo, 11.10.1970, Sili6 (SARA; isotype: LJU).

Willdenowia 38 2008 375 Note. This taxon was described as a species by Sili6 (1979) and included as a subspecies under Satureja cristata by Greuter & al. (1986). This concept is followed here, making a new combination under Micromeria necessary. M. cristata subsp. orientalis P. H. Davis in Notes Roy. Bot. Gard. Edinburgh 38: 41. 1980. Holotype: Turkey B6 Malatya: Gurun to Malatya, 40 km from Malatya, c. 1400 m, cliff ledges and cracks, fls. pale lilac, 7.8.1956, McNeill 461 (E!). M. cristata subsp. phrygia P. H. Davis in Notes Roy. Bot. Gard. Edinburgh 38: 40. 1980. Holotype: Turkey B2 Kutahya: d. Gediz, Saphane Da., 1900-2000 m, sunny rocks, flowers white with mouve spotting on lip, 27.8.1950, Davis 18457 (E!; isotype: K!). M. cristata f. subuniflora Bornm. in Bot. Jahrb. Syst. 61, Beibl. 140: 73. 1927. Holotype: [Macedonia] Macedonia centralis: In distr. Montium Duditza- et Suharupa-planina, in m. Dwe Uschi (= Zwei-Ohrenberg), c. 1700 m, 24.7.1917, Schultze 326 (B!; isotype: JE!). = M. cristata subsp. cristata M. cristata f. umbrosa Bornm. in Bot. Jahrb. Syst. 61, Beibl. 140: 73. 1927. Holotype: [Macedonia] In rupe Markov-Kamen vallis fluvii Lepenac (infra pag. Kacanik), 500 m 5.7.1918, Bornmüller, Plantae Macedonicae 4851 (HBG; isotypes: B!, BP!, JE!). = M. cristata subsp. cristata Note. Bornmüller s 1918 trip to Macedonia was financed by the Institut für Allgemeine Botanik in Hamburg, the top set of the corresponding collections, unlike other Bornmüller material, therefore is kept at HBG (H. Manitz, pers. comm.). M. cristata subsp. xylorrhiza (Boiss. & Heldr. ex Benth.) P. H. Davis in Notes Roy. Bot. Gard. Edinburgh 21: 64. 1952 M. xylorrhiza Boiss. & Heldr. ex Benth. M. croatica (Pers.) Schott in Oesterr. Bot. Wochenbl. 7: 93. 1857 Thymus croaticus Pers. in Syn. Pl. 2: 130. 1806. Type: [Croatia] Hab. in alpibus Croatiae, rupes calcareas opplens (P?). M. croatica f. glabrata (K. Maly) Sili6 in Glasn. Zemaljsk. Muz. Bosne Hercegovine Sarajevu Prir. Nauke 13: 117. 1975 Satureja croatica f. glabrata K. Maly in Glasn. Zemaljsk. Muz. Bosne Herzegovine 40: 154. 1928. Holotype: [Bosnia and Hercegovina] Lisici6i prope Konjic, solo calcareo, c. 260 m, Maly (SARA). = M. croatica (Pers.) Schott M. croatica f. linearilanceolata Sili6 in Glasn. Zemaljsk. Muz. Bosne Hercegovine Sarajevu Prir. Nauke 13: 117. 1975. Holotype: [Bosnia and Hercegovina] Kanjon Rakitnice, solo calcereo, c. 400 m, 15.10.1967, Sili6 (SARA). = M. croatica (Pers.) Schott M. croatica var. longidens Rohlena in Sitzungsber. Königl. Böhm. Ges. Wiss. Prag, Math.- Naturwiss. Cl. 1912: 103. 1913 M. croatica f. longidens (Rohlena) Hayek, Prodr. Fl. Penins. Balcan. 2: 321. 1929. Holotype: [Montenegro] Auf felsigen Ufern des Flusses Piva bei Goransko (ca. 660 m), Rohlena (PRC). = M. croatica (Pers.) Schott Note. Sili6 (1979) cites this as f. longidens Rohl., whereas originally it has been described as a variety and was reduced to the rank of a forma later (see above). M. croatica f. multicaulis Sili6 in Glasn. Zemaljsk. Muz. Bosne Hercegovine Sarajevu Prir. Nauke 13: 117. 1975. Holotype: [Bosnia and Hercegovina] in valle Drezanka pr. Dreznica, c. 150 m, solo calcareo, 4.8.1900, Maly (SARA). = M. croatica (Pers.) Schott M. croatica f. ovalifolia Sili6 in Glasn. Zemaljsk. Muz. Bosne Hercegovine Sarajevu Prir. Nauke 13: 117. 1975. Holotype: [Bosnia and Hercegovina] Idbar pr. Bozi6, c. 500 m, 7.9.1970, Sili6 (SARA). = M. croatica (Pers.) Schott M. croatica var. panciciana (Briq.) Hayek, Prodr. Fl. Penins. Balcan. 2: 321. 1929 Satureja croatica var. panciciana Briq., Lab. Alp. Marit. 3: 427. Sep 1895. Lectotype (see Note): [Serbia] Mokra Gora, 1875, Panci6 8580 (BEOU). = M. croatica (Pers.) Schott

376 Bräuchler & al.: Micromeria, a synoptical update Note. On a web page containing information on the Serbian botanist Panci6 (http://pancic. bio.bg.ac.yu/engl/index.html) a lectotypification for Satureja croatica var. panciciana Briq. is indicated without citation of the source. We were not able to trace the corresponding publication. No type material of this taxon seems to be present at G. M. cunninghamii Benth., Labiat. Gen. Spec.: 730. 1835 Mentha cunninghamii (Benth.) Benth. in Candolle, Prodr. 12: 174. 1848. M. cymuligera Boiss. & Hausskn. in Boissier, Fl. Orient. 4: 569. 1879 =? Mentha sp. Note. Micromeris cymuligera has always been regarded as isolated within the genus with respect to its annual habit and special anther structure (Boissier 1879; Bräuchler & al. 2005; Bräuchler & al. 2006; Briquet 1896; Davis 1982). Weak affinities towards sect. Micromeria (Doroszenko unpubl. Ph.D. thesis, Univ. Edinburgh 1986) have been suggested based on overall similarity rather than on shared characters. Preliminary molecular data indicate this species as disassociated from Micromeria s.str. and more closely related to the genus Mentha (Bräuchler, unpubl.). However, results were not clear enough to make a final decision. The lack of a marginal vein as revealed by anatomical studies (Bräuchler, unpubl.) favours exclusion of M. cymuligera from the genus. The perfoliate inflorescence is another feature shared with Mentha rather than with Micromeria s.str., where the bracts tend to get reduced towards the tip of the inflorescence. The ecological range from what is known (humid alpine pastures, torrent beds) is similar to that of Mentha. None of the species of Micromeria s.str. occur in such a habitat. Whether this species should be included in Mentha or placed in a monotypic genus ( Cymularia ) remains to be clarified. M. cypria Kotschy in Unger & Kotschy, Ins. Cypern: 270. 1865. Lectotype (designated here by Bräuchler): [Cyprus] In scopulorum fissures montis Pentadactylos et Buffavento 1500, 4.1862, Kotschy, Pl. Ins. Cypro 338 (W-0014293!; isolectotypes: BP!, G, JE!, K!, P!). = M. microphylla (d Urv.) Benth. Note. The second syntype cited is Kotschy 390 (K!, traced neither at W nor at WU) Um Castello della Regina, 15.4.1862. It seems that less duplicates of this collection number have been distributed, therefore Kotschy 338 is chosen as lectotype. M. cypria var. villosissima H. Lindb., Iter Cypr.: 29. 1946. Holotype: [Cyprus] Ayios Hilarion, in fissuris rupis siccissimis, 1939, Lindberg (H?). = M. microphylla (d Urv.) Benth. M. dalmatica Benth. in Candolle, Prodr. 12: 225. 1848 Clinopodium dalmaticum (Benth.) Bräuchler & Heubl in Taxon 55: 979. 2006. Note. See note under M. balcanica. M. dalmatica f. angustifrons Rohlena in Vestn. Král. Ceské Spolecn. Nauk, Tr. Mat.-Prir. 2: 11. 1933 = Clinopodium dalmaticum (Benth.) Bräuchler & Heubl in Taxon 55: 979. 2006. M. dalmatica subsp. bulgarica (Velen.) Guinea in Bot. J. Linn. Soc. 64: 381. 1971 M. dalmatica f. bulgarica (Velen.) Stoj., Stefanov & Kit., Fl. Bulg. 2: 924. 1967 M. origanifolia subsp. bulgarica Velen. = Clinopodium dalmaticum (Benth.) Bräuchler & Heubl in Taxon 55: 979. 2006. M. dalmatica f. multiflora Sili6 in Glasn. Zemaljsk. Muz. Bosne Hercegovine Sarajevu Prir. Nauke 13: 115. 1975 = Clinopodium dalmaticum (Benth.) Bräuchler & Heubl in Taxon 55: 979. 2006. M. dalmatica Fenzl in Index Seminum (Vienna) 1851: [page?]. 1851, non Benth. 1848 = Mentha pulegium L., Sp. Pl. 2: 577. 1753. Note. This name is probably a mere misinterpretation and should read M. dalmatica sensu Fenzl, non Benth. instead. It has been published later as Calamintha fenzlii Vis. in Atti Reale Ist. Veneto Sci. Lett. Arti, ser. 2, 6: 300. Mar 1855, and/or M. fenzlei Regel (see name entry below). The synonymy (= Mentha pulegium L.) as provided in Govaerts & al. (2008), seems faulty regarding material traced at W and WU, which represents Clinopodium thymifolium (Scop.) Kuntze or a closely related species. We were, however, unable to consult the reference and relevant material at PAD or LE.

Willdenowia 38 2008 377 M. danaensis Danin in Willdenowia 27: 161. 1997. Holotype: Jordan, Edom, Dana Reserve, Wadi Barra area, 2 km SE of the visitor center, in crevices of smooth-faced white sandstone outcrops, 14.5.1996, Danin 963013 (HUJ;isotype:B!). M. darwinii Benth. in Candolle, Prodr. 12: 222. 1848 Clinopodium darwinii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. darwinii var. imbricatifolia Speg. in Anales Soc. Ci. Argent. 53: 251. 1902 = Clinopodium darwinii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. darwinii var. pallida Speg. in Anales Soc. Ci. Argent. 53: 251. 1902 = Clinopodium darwinii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. darwinii var. pusilla (Phil.) Speg. in Anales Soc. Ci. Argent. 53: 250. 1902 M. pusilla Phil. = Clinopodium darwinii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. [M. darwinii var. typica Speg. in Anales Soc. Ci. Argent. 53: 250. 1902, nom. inval.] Clinopodium darwinii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. darwinii var. virescens Speg. in Anales Soc. Ci. Argent. 53: 250. 1902 = Clinopodium darwinii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. debilis Pomel in Nouv. Mat. Fl. Atl.: 122. 1874. Lectotype (designated here by Bräuchler): [Algeria] Rochers de Zouia. 1400 m, Gar Roub., jn. 1855-56, Pomel (MPU-004936*!). Note. Two sheets of M. debilis Pomel exist at MPU, one of which has been labelled holotype when the types were databased (MPU-004935*!). This is the richest type collection. Since there is no original label bearing Pomel s handwriting on that sheet, it is not considered to be suitable to serve as a lectotype. The second sheet bears two individuals each labelled with a different barcode (left hand: MPU-005512*!; right hand: MPU-004936*!) and labels written by Pomel and Maire respectively. The right hand individual is a bit richer and has more leaves on it. It therefore is chosen as lectotype here. M. debilis var. mauritii Sennen, Diagn. Nouv.: 240. 1936. Holotype: Maroc: Beni-Snassen, Taforait, rochers calcaires, vers 900 m, Mauricio, Sennen, Pl. Espagne 9532 (BC!; isotypes: MPU- 009542*!, MPU-009543*!). = M. debilis Pomel M. debilis var. villosissima Batt. & Trab. in Fl. Algérie 1: 677. Dec 1890. Holotype: [Algeria] Grands rochers des Cascades à Tlemcen (AL?). = M. debilis Pomel M. densiflora Benth., Labiat. Gen. Spec.: 375. 1834. Lectotype: (designated by Pérez 1978: 157): [Spain, Canary Islands] Hab. in ins. Canariensibus, Broussonet (K!). Note. A label written by Pérez de Paz in 1976 and attached to the holotype at K indicates this as most probably not collected on the Canary Islands but similar to some forms of M. varia Benth. For the purpose of this paper, we decided to list this name as accepted until a more detailed study on its affinities is undertaken. M. densiflora var. pitardii (Bornm.) Knoche, Vagandi Mos. [page?]. 1923 M. pitardii Bornm. = M. lepida Webb & Berthel. Note. This citation is taken from Pérez (1978), who does not list a page number. We were not able to consult the reference. M. despreauxii (Briq.) Bornm. in Repert. Spec. Nov. Regni Veg. 19: 198. 1924 Satureja despreauxii Briq. in Annuaire Conserv. Jard. Bot. Genève 2: 186. 1898. Holotype: [Spain, Canary Islands] Iles Canaries, Despreaux 321 (G). = M. helianthemifolia Webb & Berthel. Note. This name was omitted by Pérez (1978) and all other treatments so far. This might be due to its listing in a footnote of the protologue of M. kuegleri. Doroszenko (unpubl. Ph.D. thesis Univ. Edinburgh 1986) placed the basionym in synonymy to M. helianthemifolia, which is followed here.

378 Bräuchler & al.: Micromeria, a synoptical update M. diffusa Lojac., Fl. Sicul. 2(2): 216. 1907. Type: [Italy, Sicily] Monti di Palermo, Maggio, Todaro (not traced). = M. graeca (L.) Benth. ex Rchb. Note. See note under M. calaminthoides. M. dolichodonta P. H. Davis in Kew Bull. 1951: 75. 1951 Clinopodium dolichodontum (P. H. Davis) Bräuchler & Heubl in Taxon 55: 979. 2006. M. domingensis Shinners in Sida 1: 96. 1962 = Clinopodium brownei (Sw.) Kuntze, Revis. Gen. Pl. 2: 514. 1891. M. douglasii (Benth.) Benth., Labiat. Gen. Spec.: 372. 1834 Thymus douglasii Benth. in Linnaea 6: 80. 1837 Clinopodium douglasii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. echioides Lacaita ex Lojac., Fl. Sicul. 2(2): 215. 1907 = M. graeca (L.) Benth. ex Rchb. Note. Strigosa would be the variety to choose the specific lectotype from (Lojacono-Pojero, Fl. Sicul. 2(2): 215. 1907: La forma piu comune e prototipa ). Furthermore see note under M. calaminthoides. M. echioides var. angustifolia Lojac., Fl. Sicul. 2(2): 216. 1907. Type: [Italy, Sicily] Palermo, Porto Empedocle? (not traced). = M. graeca (L.) Benth. ex Rchb. [M. echioides var. gasparinii Lacaita in Lojacono-Pojero, Fl. Sicul. 2(2): 216. 1907, nom. nud.] Based on a specimen from: [Italy, Sicily] M. Gebbia non procul a Palazzo-Adriana Gasparrini 1 (not traced). M. graeca (L.) Benth. ex Rchb. M. echioides var. gossypina Lojac., Fl. Sicul. 2(2): 216. 1907. Type: [Italy, Sicily] Trapani, Marsala, Lehm. (not traced). = M. graeca (L.) Benth. ex Rchb. M. echioides var. griseocanescens Lojac., Fl. Sicul. 2(2): 215. 1907. Syntypes: [Italy, Sicily] Ad rupes elatiores, Madonie, Busambra Pan., Meretimo Tineo (not traced). = M. graeca (L.) Benth. ex Rchb. M. echioides var. humillima Lojac., Fl. Sicul. 2(2): 215. 1907. Syntypes: [Italy, Sicily] Ad rupes muscosas, Palermo M. Grifone, Tineo (not traced); Palermo, 1872, Ajuti (K). = M. graeca (L.) Benth. ex Rchb. M. echioides var. strigosa Lojac., Fl. Sicul. 2(2): 215. 1907. Syntypes: [Italy, Sicily] Ad rupes montanas, Palermo, S. Martino, Cometa, Pizzuta ed altrove (not traced). = M. graeca (L.) Benth. ex Rchb. Note. See note under M. echioides. M. ekmaniana (Epling & Alain) Borhidi in Acta Bot. Hung. 37: 79. 1992 Satureja ekmaniana Epling & Alain in Brittonia 20: 156. 1968 Clinopodium ekmanianum (Epling & Alain) Harley in Kew Bull 55: 921. 2000. M. elegans Boriss. in Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR 15: 330. 1953. Holotype: [Turkey, distr. Artvin] Prope opp. Gurshany, 30.5.1910, Nesterov 68 (LE). = M. elliptica K. Koch M. ellenbeckii (Gürke) Chiov. in Savoia-Aosta, Explor. Uabi-Uebi Scebeli: 443. 1932 Satureja ellenbeckii Gürke in Bot. Jahrb. Syst. 36: 129. 1905. Holotype: [Ethiopia. Harege region] Gara Mulata Ellenbeck 533 (B ); neotype (designated here by Ryding): Ethiopia, Harege region, Gara Mulata, Gillett 5319 (K!; isoneotype: FT!). = M. imbricata var. villosa (Elly Walther & K. H. Walther) Ryding Note. The neotype was indicated by Ryding (2007: 442) for the first time, but not accompanied by the statement designated here or an equivalent. It is therefore formally chosen here by Ryding.

Willdenowia 38 2008 379 M. elliptica K. Koch in Linnaea 21: 669. 1849. Holotype: [Turkey, A8 Çoruh] Im Hochgebirge des Gaues Pertakrek auf Urgestein, c. 5000-6000' hoch, Koch (B ). Note. Davis (1982) cited the holotype at B unseen and an isotype at G seen. Although the holotype was listed as extant in Edmondson & Lack (1977), it could not be traced at B and probably has been destroyed. The holotype of M. elliptica var. nana (or more probably a fragment thereof), however, was traced at B, whereas it was not listed in Edmondson & Lack (1977). Obviously the two taxa have been confused during compilation of the list of Koch types present at B for that publication. The identity of the (fragmentary) isotype at G is questionable since the label indicates this as M. elliptica var. nana as well and provides only cryptic evidence for the specimen being a type ( H.C.K. =? Herbarium Carolus Kochii). No original material of the typical variety could be traced at B, G, GOET, P or W. Since new collections (by T. Dirmenci and colleagues) raised doubts about infraspecific classification, we decided to leave this name untypified until a more detailed investigation has been done. M. elliptica var. nana K. Koch in Linnaea 21: 670. 1849. Holotype: [Turkey, A8 Çoruh] Im Hochgebirge des Gaues Pertakrek auf Urgestein, c. 5000-6000' hoch, Koch (B!; putative isotype: G-00087141!). = M. elliptica K. Koch Note. See preceding entry. M. elliptica var. pubescens Boiss. & Kotschy ex Boiss., Fl. Orient. 4: 571. 1879. Holotype: [Turkey] Hab. in rupibus verticalibus vallis Teng prov. Musch Armeniae 6500', 9.9.1859, Kotschy, Iter Cilic.-Kurd. 453 (G-BOIS; isotypes: B!, BP!, JE!, K!, P!, W!). = M. cristata subsp. orientalis P. H. Davis Note. The text on the labels of the isotypes slightly differs from that mentioned in the protologue: Plantae ex schistosis in alpibus prope Musch lectae; In rupestribus praeruptis ad parietes excelsas vallis Teng 6500'. M. ericifolia (Roth ex Willd.) Bornm. in Repert. Spec. Nov. Regni Veg. 19: 198. 1924, nom. rejic. prop. Thymus ericifolius Roth [Bot. Cat. 2: 50. 1802, nom. nud.] ex Willd., Enum. Pl. Horti Berol.: 624. 1809 [as ericaefolius ]. Holotype: [Spain, Canary Islands] (B-W 11017!). = M. varia Benth. Note. This name has been proposed by Pérez (1978) for rejection in favour of M. varia Benth., the formal proposal, however, so far has not been published. At GOET a Roth specimen is present, almost identical to the type found in the Willdenow herbarium of B. M. eugenioides (Griseb.) Hieron. in Bol. Acad. Nac. Ci. 4: 413. 1881 Xenopoma eugenioides Griseb. in Abh. Königl. Ges. Wiss. Göttingen 19: 237. 1874 = Clinopodium gilliesii (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. euosma (W. W. Sm.) C. Y. Wu in Acta Phytotax. Sin. 10: 229. 1965 Calamintha euosma W. W. Sm. in Notes Roy. Bot. Gard. Edinburgh 9: 89. 1916 Clinopodium euosmum (W.W.Sm.) Bräuchler & Heubl in Taxon 55: 979. 2006. [M. fasciculata Benth., Labiat. Gen. Spec.: 375. 1834, nom. nud. in notas ad M. inodora] M. graeca subsp. fruticulosa (Bertol.) Guinea Note. Bentham cites this name in his description of M. inodora without indication of author. It is most likely to be interpreted as representing a new combination, both unintended and invalid, for Satureja fasciculata Raf., which in turn he listed in synonymy to M. approximata. M. fasciculata (Raf.) Strobl in Flora 68: 370. 1885, nom. illeg. Satureja fasciculata Raf., Précis Découv. Somiol.: 39. 1814 = M. graeca subsp. fruticulosa (Bertol.) Guinea Note. Strobl listed Thymus fruticulosus Bertol., Amoen. Ital.: 101. 1819, in synonymy to his combination of S. fasciculata under Micromeria and missed on the priority of the original description Bertol. in J. Bot. Agric. 2: 76. Aug 1813, as referred to in Bertoloni (1819: 101). Strobl s name is therefore illegitimate.

380 Bräuchler & al.: Micromeria, a synoptical update M. fasciculata var. gracilis (Guss.) Strobl in Flora 68: 370. 1885, nom. illeg. Satureja fasciculata var. gracilis Guss., Fl. Sicul. Syn. 2: 92. 1844. Type: not indicated (NAP?). = M. graeca subsp. fruticulosa (Bertol.) Guinea M. fasciculata var. hirsuta (Guss.) Strobl in Flora 68: 370. 1885, nom. illeg. Satureja fasciculata var. hirsuta Guss., Fl. Sicul. Syn. 2: 92. 1844. Type: not indicated. = M. graeca subsp. fruticulosa (Bertol.) Guinea M. fenzlii Regel in Fischer & Meyer, Index Seminum (St. Petersburg) 1866: 93. 1866 [as M. fenzlei Vis. ] =? Mentha pulegium L., Sp. Pl. 2: 577. 1753. Note. See note under M. dalmatica Fenzl. Regarding synonymy as provided here, the diagnosis indicates differences to specimens traced at W and WU. Thus the plant cultivated at LE based on seeds recieved from Vienna might not be identical to the species cultivated at W and PAD by that time. A final decision is not possible without consulting further literature and material at LE and PAD. The species according to the diagnosis, however, is not part of Micromeria as circumscribed here. [M. filicaulis Schott & Kotschy in Tchihatcheff, Asie Min. Bot. 2: 131. 1860, nom. nud.] M. cremnophila subsp. anatolica P. H. Davis. Note. This name is listed as an undescribed species by Tchihatcheff. At W a specimen labelled with this name ( 441. M. filicaulis Kotschy & Schott, in monte Tauro, aestate 1836, No. 702.I, Th. Kotschy ) has been identified as M. cremnophila subsp. anatolica by P. H. Davis. M. filiformis (Aiton) Benth., Labiat. Gen. Spec.: 378. 1834 Thymus filiformis Aiton, Hort. Kew. 2: 313. 1789. Lectotype (designated here by Bräuchler): [Spain] Insulae Baleares, Jacquin (BM [Banks herbarium]). Note. According to the protologue ( nat. of the Balearic islands, introduced 1770 by Mr William Malcolm. Fl. June and July, G.H. h. ) a preserved specimen from a plant of that name cultivated in the greenhouses at K prior to the description would be the first choice for a lectotype. No such specimen, however, is present at K or BM, where many specimens from plants cultivated at Kew Gardens prior to 1800 are kept. The diagnoses in Hort. Kew. vols. 1 & 2 were largely written by Dryander, who used a manuscript left by Solander (Britten 1912; Krok 1925). In this manuscript (present at the Botany Library at BM), Thymus filiformis is stated as described from a Jacquin collection or a specimen from his herbarium (Roy Vickery, pers. comm). At BM there is a sheet in the Banks herbarium that contains three gatherings: the first labelled Herb. Helvet., i.e., a collection of (supposedly) Swiss plants, made by Dick, and purchased by Banks from Pitcairn in 1775; a second labelled Hort. Pitcairne, cultivated at Pitcairn s garden and finally the third collection Insulae Baleares. Jacquin. The original description by Solander most probably was based on this latter collection and the species later has been introduced to Kew Gardens by Malcolm. The Jacquin collection to our knowledge represents the only preserved part of the original material and therefore is designated as lectotype here. Nevertheless, the authorship for the species is to be attributed to Aiton (McNeill & al. 2006: Art. 46.7, Ex. 35). Morales (1993) includes this species under M. microphylla. Until a full revision of the species group is done, we prefer to keep both taxa separate, in case of merging, the choice would have to be for M. filiformis for reasons of priority. The status of theinfraspecifictaxaofm. filiformis and its delimitations towards M. microphylla remains to be settled. [M. filiformis var. condensata (L. Chodat) Bonafè, Fl. Mallorca 4: 56. 1980, nom. inval.] = Satureja rodriguezii var. condensata L. Chodat in Bull. Soc. Bot. Geneve, ser. 2, 15: 234. 1924. Lectotype (designated here by Bräuchler): [Spain, Balearic Islands, Mallorca,] Torre d en Pau (G-00087103 [individual on the lower right hand side]!). = M. filiformis subsp. rodriguezii Note. Bonafè does neither include the basionym nor the reference to the protologue, rendering his new combination invalid. He cites the plant as M. filiformis var. condensata L. Chod.. At G no specimen labelled with this name could be traced (L. Gautier, pers. comm.). There is, however, a collection labelled as Micromeria rodriguezii collected by R. Chodat, father of L. Chodat,

Willdenowia 38 2008 381 at Torre d en Pau on 3.4.1921. This sheet consists of several individuals (fragments of one individual?) the three right hand sided of which fit the diagnosis. Since L. Chodat explicitly states that her treatment was partially based on her father s material, this sheet most probably represents part of the original material. The right hand lower individual therefore is designated as lectotype here. M. filiformis subsp. cordata (Moris ex Bertol.) Pignatti in Giorn. Bot. Ital. 111: 52. 1977 M. cordata (Moris ex Bertol.) Moris M. filiformis (Aiton) Benth. subsp. filiformis M. filiformis var. glandulosa Sennen & Pau in Treb. Inst. Catalana Hist. Nat. 3: 193. 1917 [ M. filiformis subsp. glandulosa (Sennen & Pau) Bonafè, Fl. Mallorca 4: 56. 1980, nom. inval.]. Holotype: [Spain, Balearic Islands] Soller, fentes des rochers, jusqu a 900 m, Sennen, Pl. Espagne 2038 (BC!;isotype:MA!).=M. filiformis subsp. filiformis Note. Sennen, Pl. Espagne 2039, a number included immediately after listing of the type locality, could erroneously be regarded as syntype; it is, however, the type number of M. knochei (MA!), whose description follows in the next clearly separated paragraph in Sennen s paper. Bonafè does include the basionym, but not the reference to the protologue, rendering his new combination invalid. M. filiformis subsp. minutifolia (L. Chodat) Kerguélen in Lejeunia 120: 128. 1987 Satureja filiformis var. minutifolia L. Chodat in Bull. Soc. Bot. Genève, ser. 2, 15: 246. 1924. Type: Described from Corse without indication of collector, date or precise place. Note. No original material labelled with this name could be traced at G (L. Gautier, pers. comm.). M. filiformis subsp. rodriguezii (Freyn & Janka) Bonafè, Fl. Mallorca 4: 56. 1980 M. rodriguezii Freyn & Janka M. filiformis var. rubrifolia Sennen & Pau in Treb. Inst. Catalana Hist. Nat. 3: 193. 1917. Holotype: [Spain, Balearic Islands] Soller, murs et collines (Bianor in sched.), Sennen, Pl. Espagne 2041 (BC!;isotype:MA!).= M. filiformis subsp. filiformis. M. flacca (Nábelek) Hedge in Notes Roy. Bot. Gard. Edinburgh 25: 51. 1965 Satureja flacca Nábelek in Publ. Fac. Sci. Univ. Masaryk (Brno) 70: 44. 1926 = Clinopodium molle (Benth.) Kuntze, Revis. Gen. Pl. 2: 515. 1891. M. flagellaris Baker in J. Linn. Soc., Bot. 20: 232. 1883. Note. One of currently three accepted species endemic to Madagascar. Traditionally included in Micromeria section Micromeria, they were omitted in the last synopsis of Micromeria by Morales (1993). Morphological affinities with E African members of M. sect. Pseudomelissa were assumed previously (e.g., I. Hedge, pers. comm.) but more recent studies favour the placement in a new genus (Bräuchler, unpubl. data). The lack of a marginal leaf vein along with several other features (e.g. ecology, growth form, inflorescence) supports exclusion from Micromeria as circumscribed here. M. fontanesii K. Koch in Linnaea 6: 670. 1849 & Pomel, Nouv. Mat. Fl. Atl.: 123. 1874 Satureja filiformis Desf., Fl. Atl. 2: 8, t. 121. 1798 [non Thymus filiformis Aiton,Hort.Kew.2: 313. 1789, non M. filiformis (Aiton) Benth. 1834]. Type: [Morocco?] In fissuris rupium Atlantis, Desfontaines (P, illustrated in Desf., Fl. Atlant. 2: t. 121. 1798). M. fontanesii var. depauperata Pomel, Nouv. Mat. Fl. Atl.: 123. 1874. Holotype: [Algeria] Environs d Oran, Union du Sig (?), rochers, 1852, Durando (MPU-004938*!). = M. fontanesii K. Koch. M. fontanesii var. major Batt. & Trab., Fl. Algérie 1: 677. Dec 1890. Syntypes: [Algeria] R.R. Les 2 Cèdres (Blida), Le Chenoua (not traced, AL/MPU?). = M. fontanesii K. Koch Note. See note under M. brivesii.

382 Bräuchler & al.: Micromeria, a synoptical update M. fontanesii var. typica Batt. & Trab. Fl. Algérie 1: 677. 1890 [nom. inval.?]. Syntypes: [Morocco] Tell oranais, Dahra, Tiaret, Maroc (not traced, P?). = M. fontanesii K. Koch. Note. Despite not explicitly including the type of Micromeria fontanesii, this variety name could be regarded as not validly published. The protologue, however, includes a type citation (herb. Coss.; not the type of the species) and a diagnosis. M. forbesii Benth., Labiat. Gen. Spec.: 376. 1834. Holotype: [Cape Verde Islands] Ins. Sancti Nicolai, 1822, Forbes (K!). Note. A considerably variable species with many forms on each island of the Cape Verde archipelago. Since no clear delimitations can be drawn among the variation over the whole distribution range, the varieties published are placed in synonymy to M. forbesii here. A specimen with more detailed label data collected by Forbes is present at G (G-00018940*!) and probably represents a duplicate of the holotype at K. M. forbesii var. altitudinum Bolle in Bonplandia (Hannover) 8: 282. 1860. Holotype: [Cape Verde Islands] Hab. in ins. S. Antonii jugi excelsi Cumbre rupibus siccissimis, [11.1851], Bolle (B ). = M. forbesii Benth. Note. The holotype together with Bolle s herbarium was integrated in the general herbarium at B and destroyed in WW II. To our knowledge no isotypes exist at B or at Z/ZT where some Bolle duplicates are kept. A specimen of M. forbesii collected by Bolle in 1853 (TCD!) according to the original description might represent this variety, but indication of locality is more general, the name is not on the label or elsewhere on the specimen and flowers are absent. M. forbesii var. inodora J. A. Schmidt, Beitr. Fl. Cap. Verd. Ins.: 222. 1852. Holotype: [Cape Verde Islands] Hab. in rupibus S. Antonii, pr. Riberiram grandem, 3.1851, Schmidt (not traced). = M. forbesii Benth. Note. Unlike other Schmidt material, the type is not to kept at HBG. At B, HAL, HEID and W specimens of the typical variety cited by Schmidt were traced, but not the type of var. inodora. M. formosana C. Marquand in Hooker s Icon. Pl. 33: t. 3230. 1934 = Origanum vulgare L., Sp. Pl. 2: 590. 1753. M. forsskahlii Benth., Labiat. Gen. Spec.: 379. 1834, nom. superfl. [as forskahlei ] Thymus imbricatus Forssk. M. imbricata (Forssk.) C. Chr. var. imbricata M. frivaldszkyana (Degen) Velen. in Oesterr. Bot. Z. 49: 291. 1899 Zygis frivaldszkyana Degen in Bull. Herb. Boissier 4: 523, t. 8. Jul 1896 Clinopodium frivaldszkyanum (Degen) Bräuchler & Heubl in Taxon 55: 979. 2006. M. fruticosa (L.) Druce in Bot. Exch. Club Soc. Brit. Isles 3: 421. 1914 Melissa fruticosa L., Sp. Pl. 2: 593. 1753 Clinopodium fruticosum (L.) Kuntze, Revis. Gen. Pl. 2: 516. 1891, non Forssk., Fl. Aegypt. Arab.: 107. 1775 Clinopodium serpyllifolium subsp. fruticosum (L.) Bräuchler in Taxon 55: 980. 2006. M. fruticosa subsp. barbata P. H. Davis in Kew Bull. 1951: 77. 1951 Clinopodium serpyllifolium subsp. barbatum (P. H. Davis) Bräuchler in Taxon 55: 980. 2006. M. fruticosa subsp. brachycalyx P. H. Davis in Kew Bull. 1951: 77. 1951 Clinopodium serpyllifolium subsp. brachycalyx (P. H. Davis) Bräuchler in Taxon 55: 980. 2006. M. fruticosa subsp. giresunica P. H. Davis in Notes Roy. Bot. Gard. Edinburgh 38: 39. 1980 Clinopodium serpyllifolium subsp. giresunicum (P. H. Davis) Bräuchler in Taxon 55: 980. 2006. M. fruticosa var. italica (Huter) Fen. in Webbia 28: 375. 1973 M. marifolia var. italica Huter = Clinopodium serpyllifolium (M. Bieb.) Kuntze M. fruticosa subsp. serpyllifolia (M. Bieb.) P. H. Davis in Kew Bull. 1951: 77. 1951 Nepeta serpyllifolia M. Bieb., Fl. Taur.-Cauc. 2: 40. 1808 Clinopodium serpyllifolium (M. Bieb.) Kuntze, Revis. Gen. Pl. 2: 515. 1891.